Response of the root anatomical structure of Carex moorcroftii to habitat drought in the Western Sichuan Plateau of China.

MAIN CONCLUSION: The anatomical structures of Carex moorcroftii roots showing stronger plasticity during drought had a lower coefficient of variation in cell size in the same habitats, while those showing weaker plasticity had a higher coefficient of variation. The complementary relationship between these factors comprises the adaptation mechanism of the C. moorcroftii root to drought. To explore the effects of habitat drought on root anatomy of hygrophytic plants, this study focused on roots of C. moorcroftii. Five sample plots were set up along a soil moisture gradient in the Western Sichuan Plateau to collect experimental materials. Paraffin sectioning was used to obtain root anatomy, and one-way ANOVA, correlation analysis, linear regression analysis, and RDA ranking were applied to analyze the relationship between root anatomy and soil water content. The results showed that the root transverse section area, thickness of epidermal cells, exodermis and Casparian strips, and area of aerenchyma were significantly and positively correlated with soil moisture content (P < 0.01). The diameter of the vascular cylinder and the number and total area of vessels were significantly and negatively correlated with the soil moisture content (P < 0.01). The plasticity of the anatomical structures was strong for the diameter and area of the vascular cylinder and thickness of the Casparian strip and epidermis, while it was weak for vessel diameter and area. In addition, there was an asymmetrical relationship between the functional adaptation of root anatomical structure in different soil moisture and the variation degree of root anatomical structure in the same soil moisture. Therefore, the roots of C. moorcroftii can shorten the water transport distance from the epidermis to the vascular cylinder, increase the area of the vascular cylinder and the number of vessels, and establish a complementary relationship between the functional adaptation of root anatomical structure in different habitats and the variation degree of root anatomical structure in the same habitat to adapt to habitat drought. This study provides a scientific basis for understanding the response of plateau wetland plants to habitat changes and their ecological adaptation strategies. More scientific experimental methods should be adopted to further study the mutual coordination mechanisms of different anatomical structures during root adaptation to habitat drought for hygrophytic plants.

Influence of structural reinforcements on the twist-to-bend ratio of plant axes: a case study on Carex pendula.

During biological evolution, plants have developed a wide variety of body plans and concepts that enable them to adapt to changing environmental conditions. The trade-off between flexural and torsional rigidity is an important example of sometimes conflicting mechanical requirements, the adaptation to which can be quantified by the dimensionless twist-to-bend ratio. Our study considers the triangular flower stalk of Carex pendula, which shows the highest twist-to-bend ratios ever measured for herbaceous plant axes. For an in-depth understanding of this peak value, we have developed geometric models reflecting the 2D setting of triangular cross-sections comprised of a parenchymatous matrix with vascular bundles surrounded by an epidermis. We analysed the mathematical models (using finite elements) to measure the effect of either reinforcements of the epidermal tissue or fibre reinforcements such as collenchyma and sclerenchyma on the twist-to-bend ratio. The change from an epidermis to a covering tissue of corky periderm increases both the flexural and the torsional rigidity and decreases the twist-to-bend ratio. Furthermore, additional individual fibre reinforcement strands located in the periphery of the cross-section and embedded in a parenchymatous ground tissue lead to a strong increase of the flexural and a weaker increase of the torsional rigidity and thus resulted in a marked increase of the twist-to-bend ratio. Within the developed model, a reinforcement by 49 sclerenchyma fibre strands or 24 collenchyma fibre strands is optimal in order to achieve high twist-to-bend ratios. Dependent on the mechanical quality of the fibres, the twist-to-bend ratio of collenchyma-reinforced axes is noticeably smaller, with collenchyma having an elastic modulus that is approximately 20 times smaller than that of sclerenchyma. Based on our mathematical models, we can thus draw conclusions regarding the influence of mechanical requirements on the development of plant axis geometry, in particular the placement of reinforcements.

Why are there so many sedges? Sumatroscirpeae, a missing piece in the evolutionary puzzle of the giant genus Carex (Cyperaceae).

For over a century, the origins and mechanisms underlying the diversification of the enormous temperate genus Carex (>2100 species; Cariceae, Cyperaceae) have remained largely speculative. Characteristics such as its diverse ecology, varied biogeography, and intriguing cytology have made Carex a powerful model for studying plant evolution, but its uncertain sister-group relationships hinder its use in studies that depend on accurate ancestral state estimates and biogeographic inferences. To identify the sister to Carex, we estimated the phylogeny of all genera in the Cariceae-Dulichieae-Scirpeae clade (CDS) using three plastid and two nuclear ribosomal markers. Ancestral state reconstructions of key characters were made, and a time-calibrated tree estimated. Carex is strongly supported as sister to the rare East Asian Sumatroscirpus, sole genus of a new tribe, Sumatroscirpeae trib. nov. Believed to be unique to Carex, the perigynium (prophyllar bract enclosing a flower) is in fact a synapomorphy shared with this small tribe (∼4 species) that appeared 36 Mya. We thus suggest the initial key innovation in the remarkable diversification of Carex is not the perigynium, but could be the release of mechanical constraints on perigynia through spikelet truncation, resulting in novel adaptive morphologies. Monoecy, chromosomal change, and rapid inflorescence development enabling phenological isolation may also be involved. The tiny tribe Sumatroscirpeae will provide unprecedented insights into the inflorescence homology, evolution, diversification, and biogeographic history of its sister-group Carex, one of the world's most diverse plant lineages.

Carex diaoluoshanica (Carex sect. Lageniformes, Cyperaceae), a new species from Hainan, China.

Carex diaoluoshanica, a new species of Carex sect. Lageniformes from Hainan, China, is described and illustrated. The new species is similar to C. breviscapa but differs in having wider leaves with the leaf base gradually narrowed, 5-10 cm long and petiolelike, culms subfiliform, with only two spikes, the lateral female spikes from near the culm base.

Silicified structures affect leaf optical properties in grasses and sedge.

Silicon (Si) is an important structural element that can accumulate at high concentrations in grasses and sedges, and therefore Si structures might affect the optical properties of the leaves. To better understand the role of Si in light/leaf interactions in species rich in Si, we examined the total Si and silica phytoliths, the biochemical and morphological leaf properties, and the reflectance and transmittance spectra in grasses (Phragmites australis, Phalaris arundinacea, Molinia caerulea, Deschampsia cespitosa) and sedge (Carex elata). We show that these grasses contain >1% phytoliths per dry mass, while the sedge contains only 0.4%. The data reveal the variable leaf structures of these species and significant differences in the amount of Si and phytoliths between developing and mature leaves within each species and between grasses and sedge, with little difference seen among the grass species. Redundancy analysis shows the significant roles of the different near-surface silicified leaf structures (e.g., prickle hairs, cuticle, epidermis), phytoliths and Si contents, which explain the majority of the reflectance and transmittance spectra variability. The amount of explained variance differs between mature and developing leaves. The transmittance spectra are also significantly affected by chlorophyll a content and calcium levels in the leaf tissue.

Toward an accurate taxonomic interpretation of Carex fossil fruits (Cyperaceae): a case study in section Phacocystis in the Western Palearctic.

PREMISE OF THE STUDY: Despite growing interest in the systematics and evolution of the hyperdiverse genus Carex, few studies have focused on its evolution using an absolute time framework. This is partly due to the limited knowledge of the fossil record. However, Carex fruits are not rare in certain sediments. We analyzed carpological features of modern materials from Carex sect. Phacocystis to characterize the fossil record taxonomically. METHODS: We studied 374 achenes from modern materials (18 extant species), as well as representatives from related groups, to establish the main traits within and among species. We also studied 99 achenes from sediments of living populations to assess their modification process after decay. Additionally, we characterized 145 fossil achenes from 10 different locations (from 4-0.02 mya), whose taxonomic assignment we discuss. KEY RESULTS: Five main characters were identified for establishing morphological groups of species (epidermis morphology, achene-utricle attachment, achene base, style robustness, and pericarp section). Eleven additional characters allowed the discrimination at species level of most of the taxa. Fossil samples were assigned to two extant species and one unknown, possibly extinct species. CONCLUSIONS: The analysis of fruit characters allows the distinction of groups, even up to species level. Carpology is revealed as an accurate tool in Carex paleotaxonomy, which could allow the characterization of Carex fossil fruits and assign them to subgeneric or sectional categories, or to certain species. Our conclusions could be crucial for including a temporal framework in the study of the evolution of Carex.

Genetic variation in plant morphology contributes to the species-level structure of grassland communities.

It is becoming apparent that genetic diversity can influence the species diversity and structure of ecological communities. Here, we investigated the intraspecific trait variation responsible for this relationship. We grew 10 genotypes of the sedge Carex caryophyllea, as monocultures, under standardized conditions and measured traits related to morphology, growth, and life history. The same genotypes had been prominent in determining the structure of multispecies experimental communities, equivalent in species diversity, in which the genetic diversity of the constituent plant species had been varied in parallel. The trait measurements revealed substantial phenotypic variation among Carex genotypes, related predominantly to differences in physical size and to the spatial deployment of above- and belowground tissue. Genotypes successful in experimental communities were larger in size and tended to adopt a "guerrilla" clonal growth strategy. In general, multivariate trait summaries of genotype size (and to a lesser extent, variation along a linear discriminant axis) predicted genotype and species abundance in experimental communities. However, one genotype exhibited a large disparity in this respect. The performance of this genotype lay closer to prediction when it was growing with a highly competitive grass genotype. The strength of the relationship between genotype size and performance within communities decreased with decreasing community genetic diversity. These results indicate that intraspecific trait measurements are useful for predicting and understanding community structure. They also imply that competitive interactions between the genotypes of different species play an increased role in determining phenotype in genetically impoverished communities.

Genetic introgression as a potential to widen a species' niche: insights from alpine Carex curvula.

Understanding what causes the decreasing abundance of species at the margins of their distributions along environmental gradients has drawn considerable interest, especially because of the recent need to predict shifts in species distribution patterns in response to climatic changes. Here, we address the ecological range limit problem by focusing on the sedge, Carex curvula, a dominant plant of high-elevation grasslands in Europe, for which two ecologically differentiated but crosscompatible taxa have been described in the Alps. Our study heuristically combines an extensive phytoecological survey of alpine plant communities to set the niche attributes of each taxon and a population genetic study to assess the multilocus genotypes of 177 individuals sampled in typical and marginal habitats. We found that ecological variation strongly correlates with genetic differentiation. Our data strongly suggest that ecologically marginal populations of each taxon are mainly composed of individuals with genotypes resulting from introgressive hybridization. Conversely, no hybrids were found in typical habitats, even though the two taxa were close enough to crossbreed. Thus, our results indicate that genotype integrity is maintained in optimal habitats, whereas introgressed individuals are favored in marginal habitats. We conclude that gene flow between closely related taxa might be an important, although underestimated, mechanism shaping species distribution along gradients.